KEGG   PATHWAY: sfd02024
Entry
sfd02024                    Pathway                                
Name
Quorum sensing - Sinorhizobium fredii USDA 257
Description
Quorum sensing (QS) is a regulatory system that allows bacteria to share information about cell density and adjust gene expression accordingly. All QS bacteria produce and release chemical signal molecules called autoinducers (AIs) that increase in concentration as a function of cell density. The most commonly studied AIs belong to one of the following three categories: acylated homoserine lactones, also referred to as AI-1, used by Gram-negative bacteria; peptide signals, used by Gram-positive bacteria; and AI-2, used by both Gram-negative and Gram-positive bacteria. QS can be divided into at least 4 steps: production of AIs by the bacterial cell; release of AIs, either actively or passively, into the surrounding environment; recognition of AIs by specific receptors; and leading to changes in gene regulation once they exceed a threshold concentration. The processes controlled by QS include virulence, competence, conjugation, antibiotic production, motility, sporulation, and biofilm formation.
Class
Cellular Processes; Cellular community - prokaryotes
Pathway map
sfd02024  Quorum sensing
sfd02024

Other DBs
GO: 0009372
Organism
Sinorhizobium fredii USDA 257 [GN:sfd]
Gene
USDA257_c35010  hfq; protein Hfq [KO:K03666]
USDA257_c41250  aroH; phospho-2-dehydro-3-deoxyheptonate aldolase AroH [KO:K01626] [EC:2.5.1.54]
USDA257_c52320  putative isomerase [KO:K06998] [EC:5.3.3.17]
USDA257_c09590  kdpE; transcriptional regulatory protein KdpE [KO:K07667]
USDA257_c57940  putative transcription regulator [KO:K07782]
USDA257_c05390  putative LuxR/GerE family transcriptional regulator [KO:K18098]
USDA257_c17450  extracellular solute-binding protein family 1 [KO:K02055]
USDA257_c09950  ABC transporter, binding protein [KO:K02055]
USDA257_c12440  potD; spermidine/putrescine-binding periplasmic protein [KO:K02055]
USDA257_c45410  putative periplasmic binding ABC transporter [KO:K02055]
USDA257_c45600  ydcS; putative ABC transporter periplasmic-binding protein YdcS [KO:K02055]
USDA257_c49620  putative substrate-binding component of ABC transporter [KO:K02055]
USDA257_c09930  putative spermidine/putrescine ABC transporter, permease protein [KO:K02053]
USDA257_c12420  ABC-type transporter, integral membrane subunit [KO:K02053]
USDA257_c45430  binding-protein-dependent transport systems inner membrane component [KO:K02053]
USDA257_c45640  ydcV; inner membrane ABC transporter permease protein YdcV [KO:K02053]
USDA257_c49590  binding-protein-dependent transport systems inner membrane component [KO:K02053]
USDA257_c17490  binding-protein-dependent transport systems inner membrane component [KO:K02053 K02054]
USDA257_c09940  potH2; putrescine transport system permease protein PotH [KO:K02054]
USDA257_c12430  potB1; spermidine/putrescine transport system permease protein PotB [KO:K02054]
USDA257_c45420  transport system permease protein [KO:K02054]
USDA257_c45630  ydcU; inner membrane ABC transporter permease protein YdcU [KO:K02054]
USDA257_c49600  ABC transporter permease protein [KO:K02054]
USDA257_c17440  potA2; spermidine/putrescine import ATP-binding protein PotA [KO:K02052]
USDA257_c12410  potA1; spermidine/putrescine import ATP-binding protein PotA [KO:K02052]
USDA257_c45440  potA9; spermidine/putrescine import ATP-binding protein PotA [KO:K02052]
USDA257_c45620  ydcT; putative ABC transporter ATP-binding protein YdcT [KO:K02052]
USDA257_c49610  potA10; spermidine/putrescine import ATP-binding protein PotA [KO:K02052]
USDA257_c06430  Leu/Ile/Val-binding protein [KO:K01999]
USDA257_c09710  Leu/Ile/Val-binding protein [KO:K01999]
USDA257_c21340  braC; leucine-, isoleucine-, valine-, threonine-, and alanine-binding protein BraC [KO:K01999]
USDA257_c18360  extracellular ligand-binding receptor [KO:K01999]
USDA257_c18460  extracellular ligand-binding receptor [KO:K01999]
USDA257_c11860  Leu/Ile/Val-binding protein [KO:K01999]
USDA257_c44580  branched-chain amino acid ABC transporter, periplasmic substrate-binding protein [KO:K01999]
USDA257_c37820  putative solute-binding component of ABC transporter [KO:K01999]
USDA257_c54570  putative substrate-binding component of ABC transporter [KO:K01999]
USDA257_c49440  Leu/Ile/Val-binding protein [KO:K01999]
USDA257_c49970  extracellular ligand-binding receptor [KO:K01999]
USDA257_c09720  putative transmembrane component of ABC transporter [KO:K01997]
USDA257_c21350  putative branched-chain amino acid ABC transporter [KO:K01997]
USDA257_c08890  putative ABC-transporter permease protein [KO:K01997]
USDA257_c18340  livH1; high-affinity branched-chain amino acid transport system permease protein LivH [KO:K01997]
USDA257_c18440  livH2; high-affinity branched-chain amino acid transport system permease protein LivH [KO:K01997]
USDA257_c44590  branched-chain amino acid ABC transporter, permease protein [KO:K01997]
USDA257_c54600  putative permease component of ABC transporter [KO:K01997]
USDA257_c49490  livH3; high-affinity branched-chain amino acid transport system permease protein LivH [KO:K01997]
USDA257_c49960  natD; inner-membrane translocator [KO:K01997]
USDA257_c09730  branched-chain amino acid ABC transporter, permease protein [KO:K01998]
USDA257_c21360  putative branched-chain amino acid ABC transporter [KO:K01998]
USDA257_c18330  putative branched-chain amino acid ABC transporter, permease protein [KO:K01998]
USDA257_c18430  putative branched-chain amino acid ABC transporter, permease protein [KO:K01998]
USDA257_c44600  livM; high-affinity branched-chain amino acid transport system permease protein LivM [KO:K01998]
USDA257_c49480  braE1; high-affinity branched-chain amino acid transport system permease protein BraE [KO:K01998]
USDA257_c49950  braE2; high-affinity branched-chain amino acid transport system permease protein BraE [KO:K01998]
USDA257_c54610  ABC transporter, permease [KO:K01998]
USDA257_c21370  livG5; putative branched-chain amino acid transport ATP-binding protein LivG [KO:K01995]
USDA257_c09740  livG1; high-affinity branched-chain amino acid transport ATP-binding protein LivG [KO:K01995]
USDA257_c18350  livG3; high-affinity branched-chain amino acid transport ATP-binding protein LivG [KO:K01995]
USDA257_c18450  livG4; high-affinity branched-chain amino acid transport ATP-binding protein LivG [KO:K01995]
USDA257_c44610  braF1; high-affinity branched-chain amino acid transport ATP-binding protein BraF [KO:K01995]
USDA257_c54580  putative ATP-binding component of ABC transporter [KO:K01995]
USDA257_c49470  livG6; high-affinity branched-chain amino acid transport ATP-binding protein LivG [KO:K01995]
USDA257_c49940  braF2; high-affinity branched-chain amino acid transport ATP-binding protein BraF [KO:K01995]
USDA257_c09750  livF1; high-affinity branched-chain amino acid transport ATP-binding protein LivF [KO:K01996]
USDA257_c21380  livF5; high-affinity branched-chain amino acid transport ATP-binding protein LivF [KO:K01996]
USDA257_c18240  livF3; high-affinity branched-chain amino acid transport ATP-binding protein LivF [KO:K01996]
USDA257_c18470  livF4; high-affinity branched-chain amino acid transport ATP-binding protein LivF [KO:K01996]
USDA257_c44620  braG; high-affinity branched-chain amino acid transport ATP-binding protein BraG [KO:K01996]
USDA257_c49460  livF6; high-affinity branched-chain amino acid transport ATP-binding protein LivF [KO:K01996]
USDA257_c49930  livF7; high-affinity branched-chain amino acid transport ATP-binding protein LivF [KO:K01996]
USDA257_c54590  putative ATP-binding component of ABC transporter [KO:K01996]
USDA257_c32270  hypothetical protein [KO:K01728] [EC:4.2.2.2]
USDA257_c01130  fadD; long-chain-fatty-acid--CoA ligase FadD [KO:K01897] [EC:6.2.1.3]
USDA257_c01010  putative transcriptional regulator, Crp/Fnr family [KO:K10914]
USDA257_c23960  hypothetical protein [KO:K09823]
USDA257_c41870  zinc uptake transcriptional regulator protein, Fur family [KO:K09823]
USDA257_c09330  ribD; riboflavin biosynthesis protein RibD [KO:K11752] [EC:3.5.4.26 1.1.1.193]
USDA257_c41150  traI; acyl-homoserine-lactone synthase TraI [KO:K22968] [EC:2.3.1.184]
USDA257_c41140  transcriptional regulator NgrR, LuxR family [KO:K19733]
USDA257_c10640  oppA1; periplasmic oligopeptide-binding protein OppA [KO:K15580]
USDA257_c10650  oppB; oligopeptide transport system permease protein OppB [KO:K15581]
USDA257_c10660  oppC1; oligopeptide transport system permease protein OppC [KO:K15582]
USDA257_c10670  yejF1; putative ABC transporter ATP-binding protein YejF [KO:K10823 K15583]
USDA257_c02390  appA; oligopeptide-binding protein AppA [KO:K02035]
USDA257_c12370  ddpA1; putative D,D-dipeptide-binding periplasmic protein DdpA [KO:K02035]
USDA257_c24220  gsiB; glutathione-binding protein GsiB [KO:K02035]
USDA257_c25540  agpA2; periplasmic alpha-galactoside-binding protein AgpA [KO:K02035]
USDA257_c12660  oppA2; periplasmic oligopeptide-binding protein OppA [KO:K02035]
USDA257_c48740  ddpA2; putative D,D-dipeptide-binding periplasmic protein DdpA [KO:K02035]
USDA257_c32460  agpA3; periplasmic alpha-galactoside-binding protein AgpA [KO:K02035]
USDA257_c45530  dppA3; periplasmic dipeptide transport protein DppA [KO:K02035]
USDA257_c36880  putative binding protein [KO:K02035]
USDA257_c29880  dppA2; periplasmic dipeptide transport protein DppA [KO:K02035]
USDA257_c56120  nikA; nickel-binding periplasmic protein NikA [KO:K02035]
USDA257_c51370  dppA4; periplasmic dipeptide transport protein DppA [KO:K02035]
USDA257_c02400  appB1; oligopeptide transport system permease protein AppB [KO:K02033]
USDA257_c12360  dppB2; dipeptide transport system permease protein DppB [KO:K02033]
USDA257_c24210  putative peptide transport system permease protein [KO:K02033]
USDA257_c25550  appB3; oligopeptide transport system permease protein AppB [KO:K02033]
USDA257_c48730  dppB4; dipeptide transport system permease protein DppB [KO:K02033]
USDA257_c32470  appB4; oligopeptide transport system permease protein AppB [KO:K02033]
USDA257_c45540  gsiC2; glutathione transport system permease protein GsiC [KO:K02033]
USDA257_c29870  gsiC1; glutathione transport system permease protein GsiC [KO:K02033]
USDA257_c51340  gsiC3; glutathione transport system permease protein GsiC [KO:K02033]
USDA257_c12670  dppB3; dipeptide transport system permease protein DppB [KO:K02033]
USDA257_c56130  gsiC4; glutathione transport system permease protein GsiC [KO:K02033]
USDA257_c02410  appC1; oligopeptide transport system permease protein AppC [KO:K02034]
USDA257_c12350  ddpC1; putative D,D-dipeptide transport system permease protein DdpC [KO:K02034]
USDA257_c24200  ddpC2; putative D,D-dipeptide transport system permease protein DdpC [KO:K02034]
USDA257_c25560  appC3; oligopeptide transport system permease protein AppC [KO:K02034]
USDA257_c48720  dppC5; dipeptide transport system permease protein DppC [KO:K02034]
USDA257_c32480  appC4; oligopeptide transport system permease protein AppC [KO:K02034]
USDA257_c45550  dppC4; dipeptide transport system permease protein DppC [KO:K02034]
USDA257_c29860  dppC3; dipeptide transport system permease protein DppC [KO:K02034]
USDA257_c56140  oppC2; putative oligopeptide transport system permease protein OppC2 [KO:K02034]
USDA257_c51330  appC5; oligopeptide transport system permease protein AppC [KO:K02034]
USDA257_c12680  dppC2; dipeptide transport system permease protein DppC [KO:K02034]
USDA257_c12340  gsiA2; glutathione import ATP-binding protein GsiA [KO:K02031]
USDA257_c51360  oppD4; oligopeptide transport ATP-binding protein OppD [KO:K02031]
USDA257_c25570  gsiA3; glutathione import ATP-binding protein GsiA [KO:K02031 K02032]
USDA257_c12690  oppD1; oligopeptide transport ATP-binding protein OppD [KO:K02031]
USDA257_c45560  gsiA6; glutathione import ATP-binding protein GsiA [KO:K02031 K02032]
USDA257_c32450  gsiA5; glutathione import ATP-binding protein GsiA [KO:K02031 K02032]
USDA257_c29850  gsiA4; glutathione import ATP-binding protein GsiA [KO:K02031 K02032]
USDA257_c02380  gsiA1; glutathione import ATP-binding protein GsiA [KO:K02031 K02032]
USDA257_c56150  gsiA7; glutathione import ATP-binding protein GsiA [KO:K02031 K02032]
USDA257_c48710  oppD3; oligopeptide transport ATP-binding protein OppD [KO:K02031]
USDA257_c24190  oppD2; oligopeptide transport ATP-binding protein OppD [KO:K02031]
USDA257_c51350  appF2; oligopeptide transport ATP-binding protein AppF [KO:K02032]
USDA257_c48700  ykfD2; putative oligopeptide transport ATP-binding protein YkfD [KO:K02032]
USDA257_c12700  appF1; oligopeptide transport ATP-binding protein AppF [KO:K02032]
USDA257_c24180  ykfD1; putative oligopeptide transport ATP-binding protein YkfD [KO:K02032]
USDA257_c00780  yidC; membrane protein insertase YidC [KO:K03217]
USDA257_c01820  secDF1; protein translocase subunit SecDF [KO:K12257]
USDA257_c35790  secDF2; protein translocase subunit SecDF [KO:K12257]
USDA257_c34750  hypothetical protein [KO:K03075]
USDA257_c34100  secY; protein translocase subunit SecY [KO:K03076]
USDA257_c35780  yajC; immunogenic membrane protein YajC [KO:K03210]
USDA257_c50360  secA; protein translocase subunit SecA [KO:K03070] [EC:7.4.2.8]
USDA257_c57600  ftsY; cell division protein FtsY [KO:K03110]
USDA257_c58650  secB; protein-export protein SecB [KO:K03071]
USDA257_c57660  ffh; signal recognition particle protein Ffh [KO:K03106] [EC:3.6.5.4]
Compound
C00007  Oxygen
C00009  Orthophosphate
C00059  Sulfate
C00232  Succinate semialdehyde
C00334  4-Aminobutanoate
C00547  L-Noradrenaline
C00788  L-Adrenaline
C01019  L-Fucose
C11837  N-Butyryl-L-homoserine lactone
C11841  N-(3-Oxooctanoyl)homoserine lactone
C11844  N-Heptanoylhomoserine lactone
C11845  N-(3-Hydroxy-7-cis-tetradecenoyl)homoserine lactone
C11848  2-Heptyl-3-hydroxy-quinolone
C16421  AI-2
C16463  3',5'-Cyclic diGMP
C16640  CAI-1
C18049  N-Acyl-L-homoserine lactone
C18206  cis-11-Methyl-2-dodecenoic acid
C20643  2-Heptyl-4(1H)-quinolone
C20677  N-(4-Coumaroyl)-L-homoserine lactone
C20959  (4S)-4-Hydroxy-5-phosphooxypentane-2,3-dione
C21195  N-(3-Hydroxybutanoyl)-L-homoserine lactone
C21197  N-Hexanoyl-L-homoserine lactone
C21198  N-(3-Oxohexanoyl)-L-homoserine lactone
C21199  N-Octanoyl-L-homoserine lactone
C21200  N-3-Hydroxyoctanoyl-L-homoserine lactone
C21201  N-3-Oxo-dodecanoyl-L-homoserine lactone
C21202  cis-2-Dodecenoic acid
C21220  AIP-1
C21221  CSP
C21222  CSP
C21223  Blp
C21224  SHP2
C21225  SHP3
C21226  cCF10
C21227  iCF10
C21228  cAD1
C21229  iAD1
C21230  ComX pheromone
C21231  PhrA pentapeptide
C21232  PhrC pentapeptide
C21233  PhrE pentapeptide
C21234  PhrF pentapeptide
C21235  PhrG pentapeptide
C21236  PhrH pentapeptide
C21237  PhrK pentapeptide
C21238  NprX peptide
C21239  PapR peptide
C21241  AIP-2
C21242  AIP-3
C21243  AIP-4
C21244  AgrD peptide
C21245  AgrD1 peptide
C21246  GBAP
C21382  R-THMF
Reference
PMID:22712853
  Authors
Pereira CS, Thompson JA, Xavier KB
  Title
AI-2-mediated signalling in bacteria.
  Journal
FEMS Microbiol Rev 37:156-81 (2013)
DOI:10.1111/j.1574-6976.2012.00345.x
Reference
PMID:23965960
  Authors
Verma SC, Miyashiro T
  Title
Quorum sensing in the squid-Vibrio symbiosis.
  Journal
Int J Mol Sci 14:16386-401 (2013)
DOI:10.3390/ijms140816386
Reference
PMID:24348467
  Authors
Norsworthy AN, Visick KL
  Title
Gimme shelter: how Vibrio fischeri successfully navigates an animal's multiple environments.
  Journal
Front Microbiol 4:356 (2013)
DOI:10.3389/fmicb.2013.00356
Reference
PMID:24065108
  Authors
Papaioannou E, Utari PD, Quax WJ
  Title
Choosing an appropriate infection model to study quorum sensing inhibition in Pseudomonas infections.
  Journal
Int J Mol Sci 14:19309-40 (2013)
DOI:10.3390/ijms140919309
Reference
PMID:20738404
  Authors
Heeb S, Fletcher MP, Chhabra SR, Diggle SP, Williams P, Camara M
  Title
Quinolones: from antibiotics to autoinducers.
  Journal
FEMS Microbiol Rev 35:247-74 (2011)
DOI:10.1111/j.1574-6976.2010.00247.x
Reference
PMID:25866808
  Authors
Rasamiravaka T, Labtani Q, Duez P, El Jaziri M
  Title
The formation of biofilms by Pseudomonas aeruginosa: a review of the natural and synthetic compounds interfering with control mechanisms.
  Journal
Biomed Res Int 2015:759348 (2015)
DOI:10.1155/2015/759348
Reference
PMID:11544214
  Authors
Whiteley M, Greenberg EP
  Title
Promoter specificity elements in Pseudomonas aeruginosa quorum-sensing-controlled genes.
  Journal
J Bacteriol 183:5529-34 (2001)
DOI:10.1128/JB.183.19.5529-5534.2001
Reference
PMID:11497469
  Authors
Chin-A-Woeng TF, van den Broek D, de Voer G, van der Drift KM, Tuinman S, Thomas-Oates JE, Lugtenberg BJ, Bloemberg GV
  Title
Phenazine-1-carboxamide production in the biocontrol strain Pseudomonas chlororaphis PCL1391 is regulated by multiple factors secreted into the growth medium.
  Journal
Mol Plant Microbe Interact 14:969-79 (2001)
DOI:10.1094/MPMI.2001.14.8.969
Reference
PMID:19696934
  Authors
Hughes DT, Clarke MB, Yamamoto K, Rasko DA, Sperandio V
  Title
The QseC adrenergic signaling cascade in Enterohemorrhagic E. coli (EHEC).
  Journal
PLoS Pathog 5:e1000553 (2009)
DOI:10.1371/journal.ppat.1000553
Reference
PMID:22144490
  Authors
Njoroge J, Sperandio V
  Title
Enterohemorrhagic Escherichia coli virulence regulation by two bacterial adrenergic kinases, QseC and QseE.
  Journal
Infect Immun 80:688-703 (2012)
DOI:10.1128/IAI.05921-11
Reference
PMID:23160491
  Authors
Pacheco AR, Curtis MM, Ritchie JM, Munera D, Waldor MK, Moreira CG, Sperandio V
  Title
Fucose sensing regulates bacterial intestinal colonization.
  Journal
Nature 492:113-7 (2012)
DOI:10.1038/nature11623
Reference
PMID:22778415
  Authors
Hirakawa H, Harwood CS, Pechter KB, Schaefer AL, Greenberg EP
  Title
Antisense RNA that affects Rhodopseudomonas palustris quorum-sensing signal receptor expression.
  Journal
Proc Natl Acad Sci U S A 109:12141-6 (2012)
DOI:10.1073/pnas.1200243109
Reference
PMID:14617175
  Authors
Danino VE, Wilkinson A, Edwards A, Downie JA
  Title
Recipient-induced transfer of the symbiotic plasmid pRL1JI in Rhizobium leguminosarum bv. viciae is regulated by a quorum-sensing relay.
  Journal
Mol Microbiol 50:511-25 (2003)
DOI:10.1046/j.1365-2958.2003.03699.x
Reference
PMID:12448738
  Authors
Wisniewski-Dye F, Downie JA
  Title
Quorum-sensing in Rhizobium.
  Journal
Antonie Van Leeuwenhoek 81:397-407 (2002)
DOI:10.1023/A:1020501104051
Reference
PMID:10368158
  Authors
Rodelas B, Lithgow JK, Wisniewski-Dye F, Hardman A, Wilkinson A, Economou A, Williams P, Downie JA
  Title
Analysis of quorum-sensing-dependent control of rhizosphere-expressed (rhi) genes in Rhizobium leguminosarum bv. viciae.
  Journal
J Bacteriol 181:3816-23 (1999)
DOI:10.1128/JB.181.12.3816-3823.1999
Reference
PMID:24550924
  Authors
Lang J, Faure D
  Title
Functions and regulation of quorum-sensing in Agrobacterium tumefaciens.
  Journal
Front Plant Sci 5:14 (2014)
DOI:10.3389/fpls.2014.00014
Reference
PMID:23710277
  Authors
Planamente S, Morera S, Faure D
  Title
In planta fitness-cost of the Atu4232-regulon encoding for a selective GABA-binding sensor in Agrobacterium.
  Journal
Commun Integr Biol 6:e23692 (2013)
DOI:10.4161/cib.23692
Reference
PMID:16943991
  Authors
Barnard AM, Salmond GP
  Title
Quorum sensing in Erwinia species.
  Journal
Anal Bioanal Chem 387:415-23 (2007)
DOI:10.1007/s00216-006-0701-1
Reference
PMID:17360277
  Authors
Barnard AM, Bowden SD, Burr T, Coulthurst SJ, Monson RE, Salmond GP
  Title
Quorum sensing, virulence and secondary metabolite production in plant soft-rotting bacteria.
  Journal
Philos Trans R Soc Lond B Biol Sci 362:1165-83 (2007)
DOI:10.1098/rstb.2007.2042
Reference
PMID:19843225
  Authors
Carlier A, Burbank L, von Bodman SB
  Title
Identification and characterization of three novel EsaI/EsaR quorum-sensing controlled stewartan exopolysaccharide biosynthetic genes in Pantoea stewartii ssp. stewartii.
  Journal
Mol Microbiol 74:903-13 (2009)
DOI:10.1111/j.1365-2958.2009.06906.x
Reference
PMID:7665477
  Authors
Beck von Bodman S, Farrand SK
  Title
Capsular polysaccharide biosynthesis and pathogenicity in Erwinia stewartii require induction by an N-acylhomoserine lactone autoinducer.
  Journal
J Bacteriol 177:5000-8 (1995)
DOI:10.1128/JB.177.17.5000-5008.1995
Reference
PMID:19761612
  Authors
O'Grady EP, Viteri DF, Malott RJ, Sokol PA
  Title
Reciprocal regulation by the CepIR and CciIR quorum sensing systems in Burkholderia cenocepacia.
  Journal
BMC Genomics 10:441 (2009)
DOI:10.1186/1471-2164-10-441
Reference
PMID:16041013
  Authors
Malott RJ, Baldwin A, Mahenthiralingam E, Sokol PA
  Title
Characterization of the cciIR quorum-sensing system in Burkholderia cenocepacia.
  Journal
Infect Immun 73:4982-92 (2005)
DOI:10.1128/IAI.73.8.4982-4992.2005
Reference
PMID:22949660
  Authors
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Related
pathway
sfd02040  Flagellar assembly
KO pathway
ko02024   
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